Morphological and Systematic Interpretation of Some Late Cretaceous (Turonian-Santonian) Irregular Echinoids, Kopet-Dagh Basin, NE Iran

Home , Echinocorys, Micraster

Journal of Sciences, Islamic Republic of Iran 24(3): 243- 257 (2013) http://jsciences.ut.ac.ir University of Tehran, ISSN 1016-1104

Morphological and systematic interpretation of some Late Cretaceous (Turonian-Santonian) irregular echinoids, Kopet-Dagh Basin, NE Iran

J. Noorbakhsh Razmi1, A. A. Aryaei1, M. Taherpour Khalil Abad2* and A. R. Ashouri1&3

1 Department of Geology, Mashhad Branch, Islamic Azad University, Mashhad, Islamic Republic of ran 2 Young Researchers Club and elites, Mashhad Branch, Islamic Azad University, Mashhad, Islamic Republic of ran 3 Department of Geology, Faculty of Sciences, Ferdowsi University of Mashhad, Islamic Republic of ran

Received: 10 June 2013 / Revised: 15 July 2013 / Accepted: 19 August 2013

Abstract Abderaz Formation is one of the Upper Cretaceous formations in the Kopet- Dagh sedimentary basin. A stratigraphic section from the mentioned formation is selected for detailed systematic description with regard to echinoids. Numerous well preserved representatives of the families Mircrasteridae wright, 1857 and Echinocorythidae Lambert, 1920 are described from the Ghaleh-Zoo stratigraphic section in the Kopet-Dagh sedimentary basin, northwest of Shirvan township (NE Iran). The age of the fossil assemblage is determined as Turonian-Santonian based on four echinoid species: Micraster (Micraster) coranguinum Leske, 1778, Micraster (Micraster) cortestudinarium (Goldfuss, 1829), Echinocorys gravesi (Agassiz & Desor, 1847) and Echinocorys ex gr. scutata Leske, 1778.

Keywords: Echinoids; Late Cretaceous; Kopet-Dagh; NE Iran

proposed by geologists of the National Iranian Oil Introduction Company [2 & 3]. According to the palaeontological Irregular echinoids first appeared in the Early investigations by Kalantari (1969) in the Sheikh and Jurassic and diversified markedly during the Cretaceous Zangulalu stratigraphic sections, the age is determined and Cenozoic, attaining a near-worldwide distribution. as Turonian-Santonian. The Abderaz Formation is In the Turonian and Coniacian, the group was represented by monotonous bluish-grey to light green- dominated by members of the orders Spatangoida and grey shales. It conformably overlies the Atamir Holasteroida. The former is represented mainly by the Formation and is overlain by the Abtalkh Formation genus Micraster, the latter by Echinocorys [21]. This (figs. 2-3). In the eastern part of the Kopet-Dagh the study aims to present the results on the echinoids for the formation attains a thickness of 500 m in the southeast Abderaz Formation in the West of Kopet-Dagh (Shurijeh) and 1500 m in the northwest (Taher Abad). sedimentary basin. The formation is rich in foraminifera which indicate The Abderaz Formation (named after the village of Turonian-Santonian age. The Turonian age is Abderaz in southeastern Kopet-Dagh basin) was represented by 58 m of chalky limestones and

* Corresponding author, Tel: 0098-915-1025900, Fax: 0098-511- 8403490, E-mail: [email protected]

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Figure 1. Location of the studied section in NE Iran calcareous shales containing Ataxophtagmium puschi, Frondicularia cordata, Globotruncana helvetica, Globotruncana imbricata, Globotruncana linneiformis and Marsonella turris. As well, the Coniacian stage, 25 m thick is composed of marly limestones and calcareous shales, indicated by Gavellinella costata, Globotruncana culverensis and Globotruncana schneegansi. The grey marly limestones and black calcareous shales, about 750 m thick, are attributed to the Santonian on the basis of the following assemblage: Globotruncana concavata carinata, Globotruncana lapparenti coronate, Globotruncana lapperenti lapperenti, Globotruncana fornicata, Neoflabellina ovalis and Stensioina exculpta [29].

Figure 2. The gradual boundary between Atamir and Geological setting Abderaz formations

The NE active fold belt of Iran, Kopet-Dagh, is formed on Hercynian metamorphosed basement at the SW margin of the Turan Platform. The Kopet-Dagh region of NE Iran exposes rather complete Jurassic sections, which reach thickness of about 3000 m. The belt is composed of about 10 km thick Mesozoic and Tertiary sediments (mostly carbonates) and like the Zagros, was folded into long linear NW-SE trending folds during the last phase of the Alpine Orogeny, in the Plio-Pleistocene time. No igneous rocks are exposed in Kopet-Dagh except for those in the basement in Aghdarband tectonic window [29]. The Shirvan area is a part of the Kopet-Dagh sedimentary basin which is located in northern Khorasan province. The Ghaleh-Zoo stratigraphic Figure 3. The gradual boundary between Abderaz section is located 21 km northwest from Shirvan and Kalat formations

244 Morphological and systematic interpretation of some Late Cretaceous (Turonian-Santonian) irregular…

township (37 31’ 23’’N and 57 47’ 24’’E). The Abderaz to be taken into the every part of the body. Echinoderms and Kalat formations are well-exposed in the studied developed an internal calcareous skeleton, the so-called area. In this study the Abderaz Formation is investigated theca, which consist of fixed plates or plates of CaCO3 in the Ghaleh-Zoo stratigraphic section with regard to connected by joints. The name of this entire phylum is echinoids (Fig.1). based on the fact that there are usually numerous spines sticking through the skin and covers the calcareous skeleton to appear on the surface. Living representatives Previous studies in the Middle East of the echinoderms are subdivided into five classes and In the Kopeh-Dagh region, the first study on the of these the subphyla Blastozoa, Crinozoa and echinoderms of the Abderaz Formation (Turonian- Echinozoa are particularly important for paleontology Coniacian) was carried out by Vahidinia & Aryaei, [15]. 2000 and those of the Tirgan Formation by Hashemian The material studied comprises almost 150 et al., 2007. Likewise, a study on the Early Cretaceous specimens and have been analyzed biometrically. Most echinoderms of central Iran was carried out by of the specimens studied were collected by the author Yaghoubi et al., (2008). The latest work on the Early and are now housed at the Department of Geology, Cretaceous echinoderms in the Kopet-Dagh basin Mashhad Branch, Islamic Azad University, Mashhad, (Tirgan Formation) is carried out by Taherpour Khalil Iran and used the IAUM (Islamic Azad University, Abad et al. (2011). In this report, some specious are Mashhad Branch) as the prefix abbreviation. attributed to Toxaster (i.e. T. renevieri, T. collegnii, T. granosus), Heteraster (i.e. H. cf. delgadoi) and Loriolia.

Meanwhile, from elsewhere some species such as Heteraster musandamensis (Aptian age) and Heteraster Systematic Descriptions aff. couloni (index for Hauterivian-Barremian age) are The systematic is given following the latest concepts reported from upper Musandam limestone [14], by Smith (1984), Smith & Wright (1999, 2000, 2003) Heteraster oblongus of the Early Aptian [5], in the and descriptive terminology by Durham & Wagner United Arab Emirates. Toxaster radula, Toxaster lamberti, Toxaster dieneri and Toxaster collegnoi have been reported from the Aptian of Risan Aneiza Formation [1] in Egypt, Heteraster oblongus is reported from the Barremian of Qishn Formation [13] in Yemen; Heteraster oblongus from Sarmord Formation in North Iraq, Toxaster retusus from Turkmenistan [4], Heteratser delgadoi from the Palmyrides chain in Central Syria [1] and Heteraster renngarteni from Central Syria [35]. Figure 4. Basic measurements in the genus Micraster Agassiz, 1836; L – length of the test (in mm); W – width of the test (in mm); H – height of the test (in mm); WT – Materials and Methods width of the test measured at one third from posterior edge (in mm); AA – anal angle (in degrees); a – distance Echinoderms are marine, solitary and usually benthic from central point of apical disc to anterior edge (in mm); animals. They were diverse in shapes (in ambulacra, b – distance from central point of apical disc to posterior genital plates and etc.) in this phylum from the Early edge (in mm) (Olszewska-Nejbert, 2007). Paleozoic. Echinoderms are characterized by the presence of an ambulacral system. This organ helps the animal in food obtaining, the vascular system, the respiratory system, as well as organs for locomotion. The system starts at the surface with an opening known as the hydropore, or with a perforated calcareous madrepore plate. Water which circulates through the ambulacral system not only provides the organism with oxygen, but Figure 5. Basic measurements in the genus Echinocorys also moves microscopic particles of food towards the Leske, 1778; L – length of test; W – width of test; H – height of test; HE – height of ambitus (Olszewska- mouth. Water penetrates this to the water vascular Nejbert, 2007). system, gradually passing into the radial canal in order

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(1966) and Olszewska (2007). Also, in the systematic Biometric calculations: The characters which are paleontology studies, the biometric parameters are measured and abbreviations used for representatives of calculated in order to describe the genus and specious in this family are shown in tables 1-6. the best way. The results of these calculations are described in the biometric calculation part. Occurence: The genus Micraster first occurs in the Order SPATANGOIDA Agassiz, 1840 Cenomanian (Devon, England, e.g. Smith 1988) and Infraorder MICRASTERINA Fischer, 1966 ranges through Paleocene (Danian) of Europe [23], Family MICRASTERIDAE Lambert, 1920 Mediterranean region, Asia (Georgia, Mangyshlak and Genus Micraster Agassiz, 1836 Kopet-dagh), Madagascar, and Cuba. Micraster (Micraster) coranguinum Leske, 1778 Micraster was a deposit feeder. Test morphological (Pl. 1, figs. 1, 3; Pl. 2, Fig. 2, Pl. 3, Fig. 1; Pl. 4, figs. features show an adaptation to a very fine-grained 1, 3; Pl. 5, figs. 1-3; Pl. 7, Fig. 1) sediment [19, 27 & 28]and an infaunal mode of life. Micraster (Gibbaster) is more pyramidal in shape with 1734 Spatangus Coranguinum (a) Anglicum; J.T. subanal protofasciole, and the periproct is situated Klein, p. 33, pl. 23, figs A, B. relatively low. These characters indicate a seminfaunal 1778 Spatangus Coranguinum Var. a Anglicum; N.G. mode of life similar to recent Spatangus raschi Loven Leske, p. 221, pl. 23, figs C, D. [20]. 1829 Spatangus coranguinum Lamarck; A. Goldfuss, p. 157, pl. 48, fig. 6. Diagnostic Features: Test cordiform with anterior 1853 Micraster coranguinum Agassiz; A. D’Orbigny, sulcus; posterior face truncate. Apical disc central; p. 207, pl. 867, figs ?1-?8; pl. 868, figs ?3, ?4. ethmophract; with four gonopores. Anterior 1869 Micraster coranguinum Agassiz; G. Cotteau & J. ambulacrum in shallow sulcus from apex to peristome. Triger, p. 326, pl. 55, figs 5-10. Pore-pairs differentiated aborally; small, round with 1874 Spatangus (Micraster) coranguinum; F.A. strongly raised interporal knob. Other ambulacra Quenstedt, p. 644, pl. 87, figs 28, ?33. forming straight, shallowly sunken petals aborally; 1876 Micraster coranguinum Agassiz; G. Cotteau, p. pore-pairs elongate; perradial line grooved in type 501, pl. 83, figs 4,5. species. Peristome close to anterior margin; facing 1878 Micraster coranguinum Klein; TH. Wright, p. forward and partially or completely covered by the 271, pl. 62, figs 1-3, 5. labral plate in oral view. Labral plate longitudinally 1959 Micraster coranguinum Klein; M.M. Moskvin, elongate; sternal plates symmetic; episternal plates not p. 281, text-fig. 94; pl. 20, fig. 2. narrowing. Periproct at top of steeply truncated 1964 Micraster coranguinum (Klein); G.N. posterior face. Aboral tuberculation of uniform small Dzhabarov, p. 55, pl. 18, ?fig. 2. tubercles set in a fine groundmass of granules. Subanal 1966 Micraster coranguinum (Klein); G. Ernst, p. 124. fasciole present; ovate (Fig. 4). 1966 Micraster (Micraster) coranguinum (Leske); A.G. Fischer, p. U581, text-fig. 467,2. Remarks: This is one of the best known Cretaceous 1968 Micraster coranguinum (Klein); S.S. Mnczyska, echinoids and has formed the basis for classic p. 108, text-pl. 2, figs 1, 2; text-pl. 3, fig. 3; pl. 3, figs 1- microevolutionary studies. Stokes (1975), Ernst (1970, 3. 1972), Maczynska (1968) and Fouray (1981) deal in 1968 Micraster coranguinum (Klein); S.I. Pasternak & depth with the taxonomy of many of the Cretaceous al., p. 221, text-fig. 49; pl. 50, figs 1-4. species. Nichols (1959) made a classic analysis of the 1969 Micraster coranguinum (Klein); L. Cayeux, p. mode of life of Micraster. Micraster differs 37, pl. 1, figs 5-9. from Gibbaster in having pores in the frontal 1974 Micraster coranguinum Leske; G. Ernst & M.-G. ambulacrum that are differentiated (non-petaloid) and Schulz, p. 30, text-figs 7, 8b-c, 9d-e; pl. 1, fig. 3; pl. 2, presumably associated with funnel-building tube- fig. 2. feet. Isaster and Cyclasterhave an apical disc with three 1974 Micraster coranguinum (Klein); O.V. gonopores, whereas Micraster always has Savchinskaya, p. 328, pl. 113, figs 7-10. four. Mokotibasterdiffers in having almost no posterior 1975 Micraster coranguinum (Leske); R.B. Stokes, p. truncate face and the periproct is subambital. 64, text-fig. 29d; pl. 2, figs 1-3. Pseudogibbaster is more inflated and has a peristome 1993 Micraster coranguinum (Leske); E.P.F. Rose & that faces downwards, not forwards. N.E. Cross, text-figs 1, 5, 6. 1994 Micraster coranguinum (Leske); N.E. Cross &

246 Morphological and systematic interpretation of some Late Cretaceous (Turonian-Santonian) irregular…

Pl.1 (Scale bar 5 cm) Pl.3 (Scale bar 5 cm) 1a-c (Dorsal view, Ventral view, A): Micraster coranguinum 1a-c: Micraster coranguinum (Leske, 1778). Sample No. (Leske, 1778). Sample No. MQ116 MQ121 2a-c (Dorsal view, Ventral view, A): Micraster 2a-c: Micraster cortestudinarium (Goldfuss, 1829). Sample cortestudinarium (Goldfuss, 1829). Sample No. MQ125 No. MQ129 3a-c (Dorsal view, Ventral view, A): Micraster coranguinum 3a-c: Micraster cortestudinarium (Goldfuss, 1829). Sample (Leske, 1778). Sample No. MQ139 No. MQ152

Olszewska-Nejbert, Pl. 3, Fig. 3.

Type specious: Spatangus coranguinum (a) anglicum (Leske, 1778).

Material: From the total collected samples, 24 samples belong to this specimen.

Description: The size and thickness of the test is medium, the total shape is elliptical to heart-shape. From the side view, it is long and partly high and has a sharp dip in the anterior view. The side view is concave and the anterior groove is partly profound. The petals are shallow. The adoral side is granular in the both side of the plastron. Plastron is embossed and is covered by

Pl.2 (Scale bar 5 cm) labrum. Tubercles are small in the aboral side and are 1a-c: Micraster cortestudinarium (Goldfuss, 1829). Sample big in the adoral side. The aperture is near to the No. MQ110 anterior ridge. Periproct is circular generally. Tubercles 2a-c: Micraster coranguinum (Leske, 1778). Sample No. are covered the interambulacra. Sub-anal fasciol is MQ164 existing. The average of W/L*100 is equal to 97.68 and 3a-c: Micraster cortestudinarium (Goldfuss, 1829). Sample its median is 98.11 which is an evidence for its circular No. MQ119 form and is more close to this form rather than Micraster (Micraster) cortestudinarium. Also, the E.P. F. Rose, text-fig. 1. average of a/b*100 is equal to 105.79 and its median is 2002 Micraster coranguinum (Leske); A.B. Smith & 100 which shows that the apical system is not central C. W. Wright, p. 293, pl. 60, figs 6-8. and a little far from the center. 2007 Micraster (Micraster) coranguinum (Leske); D.

247 Vol. 24 No. 3 Summer 2013 J. Noorbakhsh Razmi et al. J. Sci. I. R. Iran

Pl.4 (Scale bar 5 cm) Pl.5 (Scale bar 5 cm) Micraster coranguinum (Leske, 1778) Micraster coranguinum (Leske, 1778) 1a-c: Sample No. MQ163 1a-c: Sample No. MQ148 2a-c: Sample No. MQ127 2a-c: Sample No. MQ118 3a-c: Sample No. MQ149 3a-c: Sample No. MQ167

Pl.6 (Scale bar 5 cm) Pl.7 (Scale bar 5 cm) Micraster cortestudinarium (Goldfuss, 1829) 1a-c: Micraster coranguinum (Leske, 1778), Sample No. MQ122 1a-c: Sample No. MQ143 2a-c: Micraster cortestudinarium (Goldfuss, 1829), Sample No. 2a-c: Sample No. MQ102 MQ151 3a-c: Sample No. MQ134 3a-c: Micraster cortestudinarium (Goldfuss, 1829), Sample No. MQ157

Remarks: Micraster (M.) coranguinum is closest to Occurrence: Late Coniacian (Magadiceramus M . (M.) bucailli but is, however, more oval in outline subquadratus Zone) at Shakh-Bogota and ?Middle and, more importantly, differs in peristome and labrum Coniacian at Sulu-Kapy. This species is widespread in features. the North European Province: Late Coniacian-

248 Morphological and systematic interpretation of some Late Cretaceous (Turonian-Santonian) irregular…

Pl.8 (Scale bar 5 cm) Pl.9 (Scale bar 5 cm) Echinocorys ex. gr. scutata (Leske, 1778) Echinocorys ex. gr. scutata (Leske, 1778) 1a-d: Sample No. EQ102 1a-d: Sample No. EQ111 2a-d: Sample No. EQ103 2a-d: Sample No. EQ110 3a-d: Sample No. EQ112 3a-d: Sample No. EQ113

Santonian of England, France, and Germany; Coniacian 1959 Micraster cortestudinarium Goldfuss; M.M. of western Ukraine and Donbass; Late Coniacian- Moskvin, p. 280, text-fig. 93; pl. 19, fig. 2; pl. 20, fig. 1. Santonian of Northern Caucasus and Kopet-Dagh; 1964 Micraster cortestudinarium (Goldfuss); G.N. Santonian of Poland (Kraków area). Known also from Dzhabarov, p. 50, pl. 17, fig. 2. the Late Coniacian- Santonian of the northern periphery 1964 Micraster carinatus G.N. Dzhabarov, p. 53, pl. of the Mediterranean Province (Georgia). 18, fig. 1. Micraster (Micraster) cortestudinarium (Goldfuss, 1967 Micraster decipiens (Bayle); L. Cayeux & O. 1829) Devilloutreys, p. 30, pl. 2, figs 7, 7a, 7b, 7c, 7B, 7Ba, (Pl. 1, Fig. 2; Pl. 2, figs. 1, 3; Pl. 3, figs. 2-3, Pl. 4, 7Bb; pl. 3, figs 7C, 7Ca. Fig. 2; Pl. 6, figs. 1-3; Pl. 7, figs. 2-3) 1968 Micraster cortestudinarium (Goldfuss); S.I. Pasternak & et al., p. 219, text-fig. 48; pl. 49, figs 5-9. 1829 Spatangus cortestudinarium A. Goldfuss, p. 156, 1970c Micraster (Micraster) cortestudinarium pl. 48, fig. 5. (Goldfuss); G. Ernst, pl. 17, fig. 3. 1874 Spatangus cortestudinarium; F.A. Quenstedt, p. 1972 Micraster cortestudinarium (Goldfuss); G. Ernst, 646, pl. 87, fig. 30. pl. 5, fig. 2. 1876 Micraster cortestudinarium Agassiz; G. Cotteau, 1974 Micraster cortestudinarium Goldfuss; O.W. p. 498, pl. 83, figs 1-3. Savchinskaya, p. 327, pl. 113, figs 1-6. 1878 Micraster cortestudinarium Goldfuss; TH. 1975 Micraster cortestudinarium Goldfuss; R.B. Wright, p. 335, pl. 76, figs 1, 2. Stokes, p. 67, text-fig. 29h. 1889 Micraster cf. cortestudinarium Goldfuss; A. Fric, 1975 Micraster decipiens (Bayle); R.B. Stokes, p. 68. p. 99, fig. 126. 1977 Micraster decipiens (Bayle); R.B. Stokes, p. 810, 1899 Micraster cortestudinarium Goldfuss; A.W. pl. 108, figs 1-4. Rowe, p. 534, pl. 35, figs iii, v. 1978 Micraster decipiens (Bayle); F. Robaszynski, pl. ?1934 Micraster cortestudinarium Goldfuss; H. Andert, 1, figs 1-4. p. 76, pl. 18, figs 19, 20. 1981 Micraster decipiens Bayle; M. Fouray, p. 38, pl.

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provenance from Quedlinburg may be excluded [34].

Material: From the total collected samples, 20 samples belong to this specimen.

Description: Generally, the size of the test is medium but some are bigger in size. The test is rather thick and generally is in heart-circular shape and rarely in circular-trapezoidal or heart-elliptical shape. The anterior side is rounded and has a rather dip groove. Petals are groove and different is length. There is a groove between the pore pairs. The shape of these pore pairs are oval to elongated oval. The aperture is located with space from the anterior ridge by an embossed ridge. The interambulacral area is covered by tubercles uniformly. Tubercles on the adoral side are bigger and compressed as well on the plastron but the ones on the adoral, are smaller. Peri-plaston plates are sandy and granular. Peristome is ovate and covered by labrum and is located farther than anterior. Periproct is circular to elliptical in shape. In some samples, the sub-anal fasciol is developed and make a closed circle. The average of

Pl.10 (Scale bar 5 cm) W/L*100 is equal to 96.86 and its median is 97.07 Echinocorys gravesi.(Agassiz and Desor,1847) which is an evidence for its circular form. Also, the 1a-d: Sample No. EQ104 average of a/b*100 is equal to 95.15 and its median is 2a-d: Sample No., EQ105 100 which shows that the apical system is not central 3a-d: Sample No., EQ106 and a little far from the center.

3, figs 8-10. Remarks: Micraster (M.) cortestudinarium differs 1984 Micraster decipiens (Bayle); B. David & M. from M. (M.) leskei and M. (M.) normanniae in being Fouray, p. 469, figs 7, 9. more circular-cordate in shape. Its maximum width is 1985 Micraster decipiens (Bayle); M. Fouray & B. located more posteriorly and its tuberculation is better Pomerol, pl. 1, figs 3, 6, 9; pl. 2, figs 2-5. developed. On the aboral side, the tubercles are larger 1991 Micraster decipiens (Bayle); R. Tarkowski, p. and more densely spaced. The type of interporiferous 130, pl. 27, fig. 1. area is “inflated”; this does not occur in M. (M.) leskei, 2002 Micraster cortestudinarium (Goldfuss); A.B. nor in M. (M.) normanniae, which also lack a Smith & C.W. Wright, p. 293, pl. 60, figs 4, 5. euamphisternal plastron. Only the semiamphisternal 2007 Micraster (Micraster) cortestudinarium (Leske); type 2 plastron appears in all three cited taxa. In such D. Olszewska-Nejbert, Text-figs 50-52; Pl. 24, Figs 1-4; cases, other characters, such as shape of test, “granular” Pl. 25, Figs 1-4; Pl. 26, Figs 1, 2. periplastronal zone, or “inflated” interporiferous area,

permit to distinguish M. (M.) cortestudinarium, which Type specious: Lectotype, designated by Stokes also has a much larger madreporite, considerably larger (1975), is the specimen illustrated by Goldfuss (1829, than other genital plates, whereas the madreporite of M. pl. 48, fig. 5 a-c) under the name of Spatangus cor (M.) leskei and M. (M.) normanniae is similar or slightly testudinarium. Possible source localities, according to larger in size to other genital plates [21]. Goldfuss (1829), are Quedlinburg, Coesfeld and Order HOLASTEROIDA Durham & Melville, 1957 Maastricht. Wood & et al. (1984) noted that the Infraorder MERIDOSTERNATA Loven, 1883 sediment preserved inside one syntype, i. e. Family ECHINOCORYTHIDAE Wright, 1857 recrystallised chalk, indicates a provenance either from Genus Echinocorys Leske, 1778 the Anglo-Paris Basin or from the chalk of Lüneburg. A

250 Morphological and systematic interpretation of some Late Cretaceous (Turonian-Santonian) irregular…

Table 1. Biometric data and simple ratios for Micraster (Micraster) cortestudinarium (Goldfuss, 1829) Number L W H WT a b W/L H/L H/W WT/W a/b of (mm) (mm) (mm) (mm) (mm) (mm) *100 *100 *100 *100 *100 specimen MQ110 50 50 20 33 25 25 100.00 40.00 40.00 66.00 100.00 MQ152 40 36 15 25 20 20 90.00 37.50 41.67 69.44 100.00 MQ132 53 52 23 38 27 27 98.11 43.40 44.23 73.08 100.00 MQ157 48 47 20 37 25 23 97.92 41.67 42.55 78.72 108.70 MQ102 51 48 22 38 27 27 94.12 43.14 45.83 79.17 100.00 MQ143 49 53 23 34 29 22 108.16 46.94 43.40 64.15 131.82 MQ125 53 51 21 39 28 26 96.23 39.62 41.18 76.47 107.69 MQ127 45 43 21 34 24 22 95.56 46.67 48.84 79.07 109.09 MQ129 39 33 19 25 20 20 84.62 48.72 57.58 75.76 100.00 MQ155 53 53 17 36 30 26 100.00 32.08 32.08 67.92 115.38 MQ134 49 52 21 37 25 25 106.12 42.86 40.38 71.15 100.00 MQ133 61 61 30 39 30 35 100.00 49.18 49.18 63.93 85.71 MQ109 51 46 26 30 27 27 90.20 50.98 56.52 65.22 100.00 MQ153 62 62 30 39 32 31 100.00 48.39 48.39 62.90 103.23 MQ137 46 43 21 30 25 23 93.48 45.65 48.84 69.77 108.70 MQ161 65 65 25 41 33 33 100.00 38.46 38.46 63.08 100.00 MQ119 52 50 18 35 30 25 96.15 34.62 36.00 70.00 120.00 MQ151 35 33 18 22 20 15 94.29 51.43 54.55 66.67 133.33 MQ146 51 47 24 29 25 25 92.16 47.06 51.06 61.70 100.00 MQ123 47 47 20 29 24 24 100.00 42.55 42.55 61.70 100.00

Table 2. Simple statistics of biometric data and simple ratios for Micraster (Micraster) cortestudinarium (Goldfuss, 1829) Statistic L W H WT a b W/L H/L H/W WT/W a/b (mm) (mm) (mm) (mm) (mm) (mm) *100 *100 *100 *100 *100 Minimum value 35.00 33.00 15.00 22.00 20.00 15.00 84.62 32.08 32.08 61.70 75.00 maximum value 65.00 65.00 30.00 41.00 33.00 35.00 108.16 51.43 57.58 79.17 116.67 Mean value 50.00 48.60 21.70 33.50 26.30 25.05 96.86 43.54 45.16 69.30 95.15 Median 50.50 49.00 21.00 34.50 26.00 25.00 97.07 43.27 43.81 68.68 100.00 Standard 7.35 8.57 3.89 5.44 3.76 4.54 5.46 5.35 6.71 5.98 9.54 Deviation

The genus Echinocorys, similar to Micraster belongs arenitic limy marls (Ernst 1970b). test height decreases to deposit feeders although it, similar to Plesiocorys, is with relative increase of clay or sand content in the epibenthonic (Ernst & Seibertz 1977; Jagt & Michels substrate. Echinicorys seems to have been much more 1994), living on soft, chalky substrate. Several sensitive to facies changes than infaunal Micraster characters, such as shape and tuberculation of the test, (compare Ernst & Seibertz 1977). large wide flattened base, the lack of the anterior groove on the ambulacral III, and of the fascioles, and the Diagnostic Features: Test ovate with flat base and character of ambulacral pores (indicating respiratory domed upper surface; no anterior sulcus. Apical disc function) on the adoral side (Smith 1980a, b, 1988; Jagt relatively large; with four gonopores. All ambulacra & Michles 1994), confirm its epibenthic mode of life. flush, with small circumflexed pore-pairs adapically. This interpretation is not commonly accepted, and Plastron meridosternous, with a single asymmetric Kongiel (1949), e. g. suggested a burrowing mode of sternal plate following the labrum. Subsequent plates life of the genus, although based on weakly evolved biserial. Periproct inframarginal to oral. No enlarged tubercles on the aboral side of the test, rare asymmetric primary tubercles aborally. No fascioles. tubercles, and the lack of fascioles in Echinocorys, Kongiel (1949) suggested it to be an ineffective, Remarks: The genus Echinocorys is rare in the shallow burrower. The shape of the test in the genus Turonian. During the Coniacian- Maastrichtian the Echinocorys is particularly sensitive for lithofacies. In number of species and individuals considerably the material from Germany, from pure limy lithotopes, increased [13 & 26]. According to the opinion of several the genus is larger and higher on average than its authors the genus Echinocorys represents a single large representatives from marly limestones, or silty to species complex in the Late Cretaceous of England.

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Table 3. Matrix correlation of biometric data and simple ratios for Micraster (Micraster) cortestudinarium (Goldfuss, 1829) Characters L W H WT A b W/L H/L H/W WT/W a/b and *100 *100 *100 *100 *100 Simple ratios L 1 0.964 0.725 0.843 0.928 0.957 0.452 -0.194 -0.350 -0.332 -0.399 W 1 0.676 0.869 0.927 0.892 0.671 -0.219 -0.457 -0.364 -0.274 H 1 0.501 0.600 0.758 0.247 0.528 0.335 -0.379 -0.415 WT 1 0.829 0.785 0.581 -0.323 -0.509 0.139 -0.264 A 1 0.824 0.535 -0.268 -0.442 -0.297 -0.071 B 1 0.328 -0.109 -0.229 -0.290 -0.604 W/L*100 1 -0.203 -0.571 -0.287 0.199 H/L*100 1 0.918 -0.138 -0.042 H/W*100 1 -0.001 -0.112 Wt/W*100 1 0.021 a/b*100 1

Table 4. Biometric data and simple ratios for Micraster (Micraster) coranguinum (Leske, 1778) Number L W H WT a b W/L H/L H/W WT/W a/b of (mm) (mm) (mm) (mm) (mm) (mm) *100 *100 *100 *100 *100 specimen MQ163 46 47 20 35 27 21 102.17 43.48 42.55 74.47 128.57 MQ121 38 39 18 27 20 19 102.63 47.37 46.15 69.23 105.26 MQ118 43 44 23 29 22 21 102.33 53.49 52.27 65.91 104.76 MQ164 48 42 23 34 25 23 87.50 47.92 54.76 80.95 108.69 MQ147 37 38 20 28 19 19 102.70 54.05 52.63 73.68 100.00 MQ105 56 56 24 35 33 30 100.00 42.86 42.86 62.50 110 MQ160 53 52 25 35 27 27 98.11 47.17 48.08 67.31 100.00 MQ103 53 52 25 33 27 26 98.11 47.17 48.08 63.46 103.84 MQ150 49 45 25 32 25 25 91.84 51.02 55.56 71.11 100.00 MQ149 43 42 22 28 22 22 97.67 51.16 52.38 66.67 100.00 MQ139 56 56 26 35 30 27 100.00 46.43 46.43 62.50 111.11 MQ116 53 51 26 33 30 25 96.23 49.06 50.98 64.71 120 MQ148 39 34 24 22 21 18 87.18 61.54 70.59 64.71 116.66 MQ101 50 49 26 28 25 25 98.00 52.00 53.06 57.14 100.00 MQ145 42 42 27 26 25 25 100.00 64.29 64.29 61.90 100.00 MQ158 49 46 29 29 25 25 93.88 59.18 63.04 63.04 100.00 MQ138 47 47 29 31 29 23 100.00 61.70 61.70 65.96 126.08 MQ167 42 42 25 25 26 26 100.00 59.52 59.52 59.52 100.00 MQ106 52 50 30 34 27 27 96.15 57.69 60.00 68.00 100.00 MQ112 48 46 26 33 25 25 95.83 54.17 56.52 71.74 100.00 MQ173 45 45 25 28 25 25 100.00 55.56 55.56 62.22 100.00 MQ122 44 44 27 31 25 24 100.00 61.36 61.36 70.45 104.16 MQ117 49 47 23 35 25 25 95.92 46.94 48.94 74.47 100.00 MQ108 50 49 30 40 27 27 98.00 60.00 61.22 81.63 100.00

Wright (1864-1882) gave an extensive list of this family are shown in Tables 7-12. synonyms of this species under the informal (nomen nudum) name Echinocorys vulgaris Breynius. However, Occurrence: Middle Turonian to Late Paleocene, Olszewska-Nejbert [21] distinguished a few varieties worldwide. within Echinocorys, and pointed out that some of the Echinocorys gravesi (Agassiz & Desor, 1847) varieties have an important stratigraphical significance, (Pl. 10, figs. 1-3) because they commonly occur within the particular lithostratigraphical units in the Late Cretaceous of 1847 Ananchytes Gravesii Desor; L. Agassiz & E. England. Desor, p. 135. 1870 Annanchytes ovata Lambert; F. Roemer, p. 312, Biometric calculations: The characters which are pl. 34, fig. 2. measured and abbreviations used for representatives of 1903 Echinocorys gravesi Desor; J. Lambert, p. 48, pl.

252 Morphological and systematic interpretation of some Late Cretaceous (Turonian-Santonian) irregular…

1, figs 12-15. 129, pl. 28, fig. 1. 1959 Echinocorys gravesi Desor; M. M. Moskvin, p. 2007 Echinocorys gravesi (Agassiz & Desor); D. 256, text-fig. 57; pl. 6, fig. 2 [=Echinocorys ex gr. Olszewska-Nejbert, Text-figs 18, 19; Pl. 8, Fig. 3; Pl. 9, scutata Leske]. Figs 1, 2; Pl. 10, Figs 1, 2. 1964 Echinocorys sphaericus (Schluter); G.N. Dzhabarov, p. 23, pl. 1, fig. 2; pl. 2, fig. 1. Type species: The holotype is specimen R 91 in the 1964 Echinocorys gravesi (Desor); G.N. Dzhabarov, collection of Agassiz & Desor, from the Craie blanche p. 25, pl. 2, fig. 2 [=Echinocorys ex gr. Scutata Leske]. of l’Oise, in the Neuchâtel Museum [21]. 1964 Echinocorys gravesi (Desor) var. moskvini; G. N. Dzhabarov, p. 26, pl. 2, fig. 3; pl. 3, fig. 1[=Echinocorys Material: From the total collected samples, 3 ex gr. scutata Leske]. samples are belong to this specimen. 1967 Echinocorys gravesi (Desor); L. Cayeux & O. De Villoutreys, p. 36, pl. 3, fig. 9 [=Echinocorys ex gr. Description: The size of the test is medium to big. It scutata Leske]. is elliptical in shape and the aboral side is high and in 1972 Echinocorys gravesi (Desor); G. Ernst, pl. 3, fig. helmet shape. The base of samples is broad and its 3; pl. 6, fig. 2. plastron is slightly concave. The side profile is concave 1974 Echinocorys sphaericus (Schluter); O.V. and is more concave in anterior side in compare with Savchinskaya, p. 321, pl. 103, figs 9-11. posterior side. The apical system is long. The pore pares 1974 Echinocorys gravesi Desor; O.V. Savchinskaya, are drop in shape and become narrow to the other pore p. 321, pl. 103, figs 12-16 [=Echinocorys ex gr. scutata pairs. Interambulacrals are few developed and are Leske]. broader than ambulacrals. Ambitus is located higher 1991 Echinocorys gravesi (Desor); R. Tarkowski, p. than its normal place. Peristome is elliptical in the

Table 5. Simple statistics of biometric data and simple ratios for Micraster (Micraster) coranguinum (Leske, 1778) Statistic L W H WT a b W/L H/L H/W WT/W a/b (mm) (mm) (mm) (mm) (mm) (mm) *100 *100 *100 *100 *100 Minimum value 37.00 34.00 18.00 22.00 19.00 18.00 87.18 42.86 42.55 57.14 100 maximum value 56.00 56.00 30.00 40.00 33.00 30.00 102.70 64.29 70.59 81.63 128.57 Mean value 47.17 46.04 24.92 31.08 25.50 24.17 97.68 53.13 54.52 67.64 105.79 Median 48.00 46.00 25.00 31.50 25.00 25.00 98.11 52.74 53.91 66.31 100 Standard Deviation 5.39 5.38 3.06 4.14 3.23 2.93 4.19 6.36 7.19 6.24 8.71

Table 6. Matrix correlation of biometric data and simple ratios for Micraster (Micraster) coranguinum (Leske, 1778) Characters L W H WT a b W/L H/L H/W WT/W a/b and *100 *100 *100 *100 *100 Simple ratios L 1 0.939 0.480 0.739 0.847 0.822 -0.137 -0.436 -0.364 -0.093 0.072 W 1 0.404 0.731 0.860 0.819 0.209 -0.458 -0.503 -0.180 0.087 H 1 0.254 0.496 0.607 -0.213 0.575 0.575 -0.171 -0.127 WT 1 0.611 0.566 0.030 -0.456 -0.441 0.537 0.097 A 1 0.800 0.051 -0.257 -0.264 -0.188 0.352 B 1 0.028 -0.129 -0.152 -0.212 -0.278 W/L*100 1 -0.100 -0.437 -0.215 0.006 H/L*100 1 0.937 -0.131 -0.183 H/W*100 1 -0.051 -0.148 Wt/W*100 1 0.038 a/b*100 1

Table 7. Biometric data and simple ratios for Echinocorys gravesi (Agassiz & Desor, 1847) Number of L W H HE W/L H/L H/W HE/H specimen (mm) (mm) (mm) (mm) *100 *100 *100 *100 EQ105 73.00 61.00 51.00 28.00 83.56 69.86 83.61 54.90 EQ106 66.00 59.00 48.00 23.00 89.39 72.73 81.36 47.92 EQ104 69.00 56.00 47.00 22.00 81.16 68.12 83.93 46.81

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anterior side and is less groove. Periproct is located in of Kopet-Dagh and Donbass, respectively, as the ridge side and is generally rounded. The apical Echinocorys sphaericus. According to Ernst (1972), system is slightly flattened. Tubercles are too small and these specimens appear to be early forms of E. gravesi. rarely can be finding in the aboral side but in the adoral side are bigger and visible. The ratio of HE/H*100 is Occurrence: Late Turonian (Inoceramus perplexus >30. Zone) of Odra I and II, Bolko, and Folwark and coeval levels in Wolin. Also known from the Middle Turonian Remarks: Echinocorys gravesi appears in the to Early Coniacian of Germany (Lower Saxony, Middle Turonian and becomes commoner in the Late Westphalia), and Late Turonian to Early Coniacian of Turonian. Some forms, particulary earlier morphotypes England, France, Donbass, Kopet-Dagh, plus Early of E. gravesi, are similar in shape to Crassiholaster Coniacian of northern Spain. sphaericus (Schluter); these occur in the Inoceramus Echinocorys ex gr. scutata Leske, 1778 (pl. 8-9) perplexus Zone (early Late Turonian) in the Opole area. Forms found in higher up section (Mytiloides scupini 1778 Echinocorys scutatus N.G. Leske, p. 111, pl. 15, Zone) in this area more closely resemble E. gravesi as figs. A, B. reported by Lambert (1903, p. 48, pl. 1, figs 12-15). 1881 Echinocorys vulgaris Breynius, T. Wright According to the report by Olszewska-Nejbert (2007), (partly), p. 328, pl. 77, figs 1-11. typical representatives of E. gravesi are recorded from 1903 Echinocorys vulgaris var. scutatus Leske; J. the Late Turonian of Wolin and northern France (Pl. 10, Lambert, p. 58. Figs 1, 2). Dzhabarov (1964, p. 23, pl. 1, fig. 2; pl. 2, 1959 Echinocorys gibbus Lamarck; M.M. Moskvin, p. fig. 1) and Savchinskaya (1974, p. 321, pl. 103, figs 9- 256, text-fig. 56; pl. 6, fig. 1. 11) described convex specimens from the Late Turonian 1959 Echinocorys gravesi Desor; M.M. Moskvin, p.

Table 8. Simple statistics of biometric data and simple ratios for Echinocorys gravesi (Agassiz & Desor, 1847) Statistic L W H HE W/L H/L H/W HE/H (mm) (mm) (mm) (mm) *100 *100 *100 *100 Minimum value 66.00 56.00 47.00 22.00 81.16 68.12 81.36 46.81 Maximum value 73.00 61.00 51.00 28.00 89.39 72.73 83.93 54.90 Mean value 69.33 58.67 48.67 24.33 84.71 70.24 82.96 49.88 Median 69.00 59.00 48.00 23.00 83.56 69.86 83.61 47.92 Standard Deviation 3.511885 2.516611 2.081666 3.21455 4.234652 2.328111 1.401635 4.387981

Table 9. Matrix correlation of biometric data and simple ratios for Echinocorys gravesi (Agassiz & Desor, 1847) Characters and L W H HE W/L H/L H/W HE/H sample ratios *100 *100 *100 *100 L 1 0.471 0.775 0.827 -0.627 -0.548 0.751 0.843 W 1 0.923 0.886 0.392 0.479 -0.228 0.872 H 1 0.996 0.007 0.103 0.165 0.993 HE 1 -0.080 0.017 0.250 1.000 W/L*100 1 0.995 -0.985 -0.109 H/L*100 1 -0.964 -0.012 H/W*100 1 0.278 HE/H*100 1

Table 10. Biometric data and simple ratios for Echinocorys ex gr. scutata (Leske, 1778) Number of L W H HE W/L H/L H/W HE/H specimen (mm) (mm) (mm) (mm) *100 *100 *100 *100 EQ102 79.00 66.00 54.00 22.00 83.54 68.35 81.82 40.74 EQ103 70.00 61.00 50.00 20.00 87.14 71.43 81.97 40.00 EQ112 57.00 49.00 38.00 17.00 85.96 66.67 77.55 44.74 EQ113 47.00 43.00 35.00 15.00 91.49 74.47 81.40 42.86 EQ115 54.00 45.00 38.00 20.00 83.33 70.37 84.44 52.63 EQ111 53.00 46.00 39.00 13.00 86.79 73.58 84.78 33.33 EQ110 58.00 45.00 39.00 23.00 77.59 67.24 86.67 58.97 EQ116 41.00 37.00 28.00 13.00 90.24 68.29 75.68 46.43

254 Morphological and systematic interpretation of some Late Cretaceous (Turonian-Santonian) irregular…

256, text-fig. 57; pl. 6, fig. 2. 1964 Echinocorys gravesi (Desor); G.N. Dzhabarov, Material: From the total collected samples, 8 p. 25, pl. 2, fig. 2. samples belong to this specimen. 1964 Echinocorys gravesi (Desor) var. moskvini; G.N. Dzhabarov, p. 26, pl. 2, fig. 3; pl. 3, fig. 1. Description: The size of the test is various in 1966 Echinocorys scutatus Leske; C.D. Wagner & samples. The shape is generally elliptical to sub J.W. Durham, p. U528, fig. 416,8. elliptical and is completely rounded and concave in the 1967 Echinocorys gravesi (Desor); L. Cayeux & O. anterior side but is less concave in the posterior side. De Villoutreys, p. 36, pl. 3, fig. 9. Periproct is located in the ridge side and is generally 1968 Echinocorys cf. conicus Agassiz var. minor rounded. Peristome is circular to elliptical in shape. In Lambert; S.I. Pasternak & et al., p. 212, text-fig. 42; pl. some samples, the regular tubercles are visible from the 44, figs 6, 7. aperture to the ridge side. There are no tubercles on the 1970 Echinocorys scutata Leske; N.B. Peake & R.V. aboral side or rarely are founded. The apical system is Melville, p. 57, pl. 2, figs A, B. long. There is no labrum and fasciol. Ambitus is located 1974 Echinocorys gravesi Desor; O.V. Savchinskaya, higher than its normal place. The ratio of HE/H*100 is p. 321, pl. 103, figs 12-16. >30. 1974 Echinocorys ex gr. scutata Leske; G. Ernst & Remarks: Leske (1778) was the first to use the M.G. Schulz, p. 36, text-figs 12, 13; pl. 4, figs 1-4. binomen Echinocorys scutatus, with masculine ending. 2002 Echinocorys scutata Leske; A.B. Smith & C.W. Hayward (1940) corrected it to “scutata”. Ernst & Wright (pars), p. 287, text-fig. 13.1(A-D, I-J, K-L, O-P); Schulz (1974) noted a wide range of variation in pl. 59, figs 1, 2. Echinocorys ex gr. scutata, first occurring in the Middle 2003 Echinocorys scutata Leske; A.B. Smith & C.W. Coniacian. They distinguished several morphotypes and Wright (pars), p. 531, text-fig. 218; pl. 168, figs 1-4, pl. some of them regarded as formal subspecies (e.g. 169, fig. 5; pl. 170, figs 1-3, 8-9; pl. 171, figs ?1-?3, 4- Echinocorys scutata scutata Leske, or E. scutata 9. vulgaris Breynius), the others as morphotypes (e.g. 2007 Echinocorys ex gr. scutata Leske; D. Olszewska- “planodoma”) only. Smith & Wright (2003) enclosed all Nejbert, Text-figs 18, 19; Pl. 8, Fig. 3; Pl. 9, Figs 1, 2; specimens of Echinocorys from the Late Cretaceous of Pl. 10, Figs 1, 2. England under the name Echinocorys scutata and distinguished 11 informal forms (scutata, planodoma Type species: Neotype figured by Peake & Melville with extreme forms named depressula, elevata, (1970, pl. 2, figs A, B) is specimen NHM E.8721, from gravesii, cincta, pyramidalis, ovata, subglobosa, conica, Fletcher’s Pit, Gravesend, Kent, England; upper vulgaris, depressa). Micraster coranguinum Zone, Early Santonian. Some of the forms illustrated by Smith & Wright

Table 11. Simple statistics of biometric data and simple ratios for Echinocorys ex gr. scutata (Leske, 1778) Statistic L W H HE W/L H/L H/W HE/H (mm) (mm) (mm) (mm) *100 *100 *100 *100 Minimum value 41.00 37.00 28.00 13.00 77.59 66.67 75.68 33.33 Maximum value 79.00 66.00 54.00 23.00 91.49 74.47 86.67 58.97 Mean value 57.38 49.00 40.13 17.88 85.76 70.05 81.79 44.96 Median 55.50 45.50 38.50 18.50 86.38 69.36 81.89 43.80 Standard Deviation 12.15 9.67 8.24 3.94 4.37 2.91 3.69 7.92

Table 12. Matrix correlation of biometric data and simple ratios for Echinocorys ex gr. scutata (Leske, 1778) Characters and L W H HE W/L H/L H/W HE/H sample ratios *100 *100 *100 *100 L 1 0.975 0.983 0.725 -0.444 -0.215 0.286 -0.181 W 1 0.982 0.588 -0.237 -0.093 0.157 -0.351 H 1 0.656 -0.350 -0.033 0.342 -0.291 HE 1 -0.803 -0.450 0.502 0.530 W/L*100 1 0.542 -0.652 -0.612 H/L*100 1 0.283 -0.550 H/W*100 1 0.223 HE/H*100 1

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(2003) are similar to morphotypes distinguished in this paper e.g. form vulgaris (Smith & Wright 2003, pl. 170, Acknowledgements figs 1, 3, 8); the other forms are different. E. scutata form depressula extreme form of planodoma (Smith & We thank the British museum for information Wright 2003, pl. 169, fig. 5) is exactly similar to E. presented on the Echinoderms. We thank Dr. Andrew scutata morphotype “vulgaris-planodoma” (Pl. 14, figs Smith, Department of paleontology, the Natural History 1-4) and is nearly similar to E. ex gr. scutata type Museum for his helpful guides and review on the vulgaris of Ernst & Schulz (1974, pl. 4, fig. 3). E. determined taxa. Reviews of the English by Dr. Ivana scutata form subglobosa [26] is exactly similar to Carevic (University of Belgrade) is gratefully “scutata” morphotype (Pl. 13, figs 1-3) and resembles acknowledged. E. ex gr. scutata type scutata of Ernst & Schulz (1974, fig. 12.1). E. scutata form ovata (Smith & Wright 2003, References pl. 171, figs 7-9) seems to be similar to “vulgaris- scutata” morphotype (Pl. 15, figs 1, 2). Moreover, it is 1. Abdelhamid M.A. and Azab M.M. Aptian-Cenomanian remarkable that E. scutata form planodoma by Smith & Echinoids from Egypt. Rev. Paleobiol., Geneve, 22(2): Wright (2003, pl. 169, figs 1-4) is different from the 851-876 (2003). same form by Ernst & Schulz (1974, fig. 12.6- 7, pl. 4, 2. Afshar-Harb A. Geology of Kopet Dagh. GSI pub., figs 1, 2) and also by Olszewska-Nejbert (2007) in (Pl. Tehran (in Persian) (1994). 3. Afshar-Harb A. History of oil exploration and brief 12, figs 1-3). description of the geology of the Sarakhs area and the anticline of Khangiran. Bull. Ir. Petrol. Ins. (BIPI), 37: Occurrence: Early Coniacian (Cremnoceramus c. 86-94 (in Persian) (1969). crassus Zone) at Shakh-Bogota, Middle and Late 4. Cecca F., Dhondt A.V., and Bogdanova T.N. The Aptian Coniacian at Sulu-Kapy, Shyrkala-Airakty and stratigraphy of the southern Tuarkyr (NW Turkmenistan, condensed Late Turonian-Coniacian of Azhirektoy and Central Asia). Riv. Ital. Paleontolo. S., 105(3): 377-396 (1999). Besakty, plus Early Coniacian at Folwark. This species 5. Dunnington H.V., Wetzel R. and Morton D.M. Iraq, has also been reported from the Middle-Late Coniacian Mesozoic and Palaeozoic, Lexique stratigraphique and Santonian of the North European Province international, ASIE, Paris, 10: 333 pp. (1959). (England, France, Belgium, Germany, western Ukraine, 6. Durham J.W. and Wagner C.D. Glossary of Donbass, ?northern Caucasus, Kopet-Dagh). morphological terms applied to Echinoids. In: R.C. Ernst & Schulz (1974) described a very similar Moore (Ed.), Treat. Invert. Paleontolo. GSA and the Echinocorys ex gr. scutata fauna from the Middle University of Kansas; Boulder, Colorado, Part U, Echinodermata 3, 1: 251-256 (1966). Coniacian to Middle Santonian from Lägerdorf, 7. Ernst G. and Seibertz E. Concepts and methods of northern Germany, with an acme in the Late Coniacian Echinoid Biostratigraphy. In: E.G. K Auffman & J.E. and Early Santonian. At Lägerdorf, the morphotype Hazel (Eds), Con. Biostrati., 541-563 (1977). “striata” succeeds the morphotype “vulgaris” during the 8. Ernst G. Grundfragen der Stammesgeschichte bei Middle Santonian. irregularen Echiniden der nordwesteuropaischen Oberkreide. Geologie. Jahrb., BGR (A), 4: 63-175 (1972). 9. Ernst G. Zur Stammesgeschichte und stratigraphischen Results and Discussion Bedeutung der Echiniden-Gattung Micraster in der nordwestdeutschen Oberkreide. Mitte. Geologi. Echinoids faunal studies in the study area led to Palaontolo., Institut der Universitat Hamburg, 39: 117- determination of 2 genus and 4 species. The recognized 135 (1970). genus are Micraster and Echinocorys. Four species have 10. Fouray M. L'evolution des Micraster (Echinides; been determined: Micraster coranginum, M. Spatangoides) dans le Touronien-Coniacien de Picardie Occidentale (Somme), Interet-biostratigraphique. Annal. cortestodinarium, Echinocorys gravesi and E. scutata. Paleontolo., Invertebres, 67: 81-134 (1981). Descriptions and standard biometric parameters are 11. Goldfuss A. Petrefacta Germaniae. Abbild-ungen und done and calculated accurately by the statistic relations Beschreibungen der Petrefacten Deutsch-land und der and various parametric data following the related charts angrenzenden Länder. Arnz & Co., Düsseldorf, 1: 1-252 and diagrams. Some essential data such as average ratio, (1826-33). minimum and maximum value, mean value, median, 12. Hashemian N. and Aryaei, A.A. Introduction of some species from Tirgan Formation, 1st Symposium of PSI, standard deviation and matrix correlation for each taxon 154-158 (2007). are reported. According to the determined echinoids, the 13. Howarth M.K. and Morris N.J. The Jurassic and Lower Turonian-Santonian age is suggested for the Abderaz Cretaceous of Wadi Hajar, southern Yemen, Bull. NHM Formation in the studied area. (Geology), 54(1): 1-32 (1998).

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